EPHEMEROPTERA, PLECOPTERA, AND TRICHOPTERA FROM SOUTHWESTERN
Transcript of EPHEMEROPTERA, PLECOPTERA, AND TRICHOPTERA FROM SOUTHWESTERN
PROC. ENTOMOL. SOC. WASH.109(1), 2007, pp. 136-154
SUMMER EPHEMEROPTERA, PLECOPTERA, AND TRICHOPTERA FROMSOUTHWESTERN DRAINAGES IN GREAT SMOKY MOUNTAINS
NATIONAL PARK, WITH ADDITIONAL EPHEMEROPTERA RECORDS
R. Edward DeWalt, Luke M. Jacobus, and W. P. McCafferty
(RED) Illinois Natural History Survey, 1916 S Oak St., Champaign, IL 61820,
U.S.A. (e-mail: [email protected]); (LMJ, WPM) Department of Entomology,
Purdue University, West Lafayette, IN 47907-1158, U.S.A.
Abstract.—Ephemeroptera (mayflies), Plecoptera (stoneflies), and Trichoptera
(caddisflies) (in total, EPT) were inventoried during May 2003, June 2003, and early
July 2004 at 15 stream reaches in southwestern Great Smoky Mountains National
Park (GRSM). Adults and immatures were collected using ultraviolet Hght traps,
sweepnetting, and handpicking. At least 169 species were collected, including 54
mayfly species, 38 stonefly species, and 77 caddisfly species. Two small, western, low
elevation streams. Shop Creek and Tabcat Creek, produced the greatest number of
species at 61 and 56 species, respectively. The two large streams. Eagle and Deepcreeks, also produced high species richness at 53 species each. A species accumulation
curve and two richness estimates demonstrated that EPT richness in this particular
area may be 80 to 120 species higher than observed. A cluster analysis showed that
adjacent stream reaches had relatively similar faunas and that different habitats
supported quite different assemblages. Significant regional records included
Ameletiis tevtiiis McDunnough and Epeorus fragilis (Morgan). Several Ephemer-
optera species are reported for the first time from North Carolina (4), Tennessee (6),
and GRSM (7). Four Trichoptera species are reported from GRSM for the first time.
Key Words: Ephemeroptera, Plecoptera, Trichoptera, Great Smoky MountainsNational Park
The National Park Service has been and McCafferty 2006). The southern
conducting an All Taxa Biodiversity Appalachian Mountain Range and
Inventory (ATBI) in the Great Smoky GRSM, in particular, are important
Mountains National Park (GRSM) since sources of Ephemeroptera (mayflies),
1997 (Sharkey 2001), one of relatively Plecoptera (stoneflies), and Trichoptera
few Park Service properties receiving any (caddisflies) (hereafter referred to as
such attention (Baumgardner and EPT) diversity (Morse et al. 1993;
Bowles 2005). The inventory efforts in Jacobus and McCafferty 2003, 2005).
GRSM have yielded data that augment DeWalt and Heinold (2005), working in
our knowledge of the diversity and the western Abrams Creek drainage of
distribution of aquatic insects in North GRSM, recorded 39 EPT species not
America, including the discovery of previously reported from GRSM, eight
several new species (Parker 2000; Peter- of which were new Tennessee state
sen et al. 2004; Etnier et al. 2004; Jacobus records. Other GRSM areas that appear
VOLUME 109, NUMBER 1 137
Table 1. Location information, approximate stream width, elevation, dates of collection, and methodsused at 15 stream sites in southwestern Great Smoky Mountains National Park in May and June 2003 andJuly 2004. Stream sites oriented from west to east.
138 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
beginning at sunset) and reflective sheet
size (1 m2). Trays of 80% EtOH were
positioned below the sheet to capture
falling insects. Mayfly subimagoes were
captured from the sheet and reared to
imagoes. Males of Perlidae and Perlodi-
dae stoneflies were captured and their
intromittent organ extruded to facilitate
species identification. Remoteness of
access meant that we could not control
for variability of weather, hence, somedifference in effort is acknowledged.
Sweepnetting of riparian vegetation for
adults and handpicking in streams for
nymphs, larvae, and pupae was also
conducted until no new species were
detected. Geographic coordinate data
were captured using a GarminTM 12XLglobal positioning system at each site.
Samples were sorted in their entirety
and specimens identified to species.
Often only the males of species could
be identified, but where it seemed that
color pattern, size, or wing veination wasconsistent with males of known identity,
they too were determined. All specimens
are housed in the INHS insect collection
and were entered into the INHS insect
collection database (INHS 2005).
An EPT species richness data matrix
was constructed with the number of
collections, number of methods, eleva-
tion, and stream wetted width (size) as
potential correlative variables. Relation-
ships among these were tested using
Spearman Rank Correlation (Cody andSmith 1991). Another matrix, one of
stream reach-by-species presence or ab-
sence was built in order to determine
efficacy of sampling. Here, all methodsfor a reach were pooled, taxa at the
genus level discarded unless they wereunique, and data from John's Coveexcluded due to poor sampling. The data
matrix was analyzed using PCOrdTMsoftware (McCune and Mefford 1999),
with species accumulation curves andjackknifing options chosen. The species
accumulation curve assesses sample ad-
equacy by repeatedly sampling the data
matrix to provide a prediction of rich-
ness versus subsample units. It also
calculates standard deviation about this
predicted value. Curves that climb rap-
idly and level out demonstrate saturation
of effort in the area. Those whose slope is
steep have not nearly exhausted the
available species.
PCOrd also provides two jackknife
estimates of species richness from ran-
dom permutations of the data matrix.
McCune and Mefford (1999) provide
a brief explanation of two procedures,
the first-order and second-order jack-
knife estimators. The first-order estima-
tor makes use of the number of species
occurring in just one sample unit (sin-
gletons), the total number of species, andthe number of sample units (Palmer
1990). The second-order estimator uses
the number of species that occur in twosample units, in addition to the single-
tons. Biases of both methods are dis-
cussed by Hellman and Fowler (1999).
Another data matrix was constructed
for species abundances by stream reach.
This data set was used in an unweighted
pair-group method cluster analysis pro-
vided in PCOrd. Options chosen includ-
ed flexible beta linkage (P=— 0.25) andthe distance measure used was Soren-
sen's (1948) similarity subtracted fromone. Five groupings were specified for
the analysis, a choice that minimized the
percent chaining of small groups to large
ones, an outcome that biases cluster
analysis results (McCune and Mefford
1999). We assumed here that adjacent
streams, those of similar size and gradi-
ent, and of the same habitat type should
cluster closer together than those that donot share these habitat similarities.
Results
Our efforts yielded 169 EPT species,
with mayflies, stoneflies, and caddisflies
contributing 54, 38, and 77 species,
respectively (Table 2). Mayflies were
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^\^^^/^^y<////^VFig. 1. Ephemeroptera, Plecoptera, and Trichoptera species richness from stream reaches in
southwestern Great Smoky Mountains National Park, Tennessee and North Carolina.
dominated by Heptageniidae (23 species
among six genera) and Ephemerellidae
(11 species among four genera). Plecop-
tera were dominated by Perlodidae (10
species among six genera) and Chloro-
perlidae (seven species among four gen-
era). Among the Trichoptera, the Hy-dropsychidae were dominant ( 1 1 species
among five genera). Nine species were
collected from each of three other
families: Leptoceridae, Polycentropodi-
dae, and Rhyacophilidae. Caddisflies
dominated species richness at 12 of 15
locations. This appeared to be a natural
pattern for reaches with diverse faunas
and wetted widths greater than 2 m.
Proctor Branch, a moderately diverse
(34 EPT) and narrow stream (2 m), wasthe only site where stoneflies contributed
the greatest number of EPT species.
Our data suggest that the most impor-
tant factor explaining species richness in
this study is the number of collections,
essentially the number of visits, to a given
reach (Spearman Rank correlation,
R=0.70, p= 0.004). Neither elevation,
wetted width, nor the number of meth-
ods used produced a significant correla-
tion. Species richness ranged from five
(Johns Creek, incompletely sampled) to
61 (Shop Creek) (Fig. 1). Five streams
could be viewed as hyperdiverse (>40species) in light of the richness of the
other 10 reaches. These included ShopCreek and Tabcat Creek, TwentymileCreek at campground 93, Eagle Creek at
campground 89, and Deep Creek.
Shop Creek and Tabcat Creek each
drained relatively open, hardwood for-
ests, had narrow widths (4—5 m wetted
width), and were at the lowest elevations
(268-299 m) of all reaches sampled.
Additionally, the Tabcat Creek drainage
was unique in that it had a well-developed
floodplain above and below the sample
reach. The remaining hyperdiverse
reaches were moderate to large streams
with high gradients. Deep Creek was the
most open canopied of the large streams.
It appears that our sampling was not
so extensive as to exhaust the area
species pool. This is evident by the slope
of the species accumulation curve being
nearly 45, with only a hint of leveling off.
Jackknife estimator procedures both
produced estimates much greater than
our current 169 species. The first-order
jackknife was 245 species, while the
second-order was 293. Between these
two estimates are 76-124 addition EPTspecies to be found in the local area of
SW GRSM.
148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
The cluster analysis produced several
clusters of stream reaches (Fig. 3). Thetwo Twentymile Creek reaches were the
most similar pair, being only three to
four miles apart and flowing through the
same elevational gradient and vegetation
type. Eagle Creek at campground 89 andEkaneetlee Creek also clustered closely.
These were adjacent, with the latter
being tributary to the former. Other
reach pairs that clustered tightly together
were Proctor Branch + Gunna Creek andTabcat Creek + Shop Creek. Cattail
Branch and its seeps also formed a clus-
ter, but at a much greater distance. Bothdrain low gradient floodplain habitat
and were curiously depauperate. Overall,
the cluster analysis suggested that manyof these reaches have great turnover in
EPT community composition and that
EFT fauna are not evenly distributed in
the region.
Significant Records
Ephemeroptera
Ameletidae
Ameletus tertius McDunnough.—Twolarvae were collected from Deep Creek.
Zloty (1996) reported A. tertius fromsoutheastern Canada, Maine, Vermont,and New York in his review of NorthAmerican species of the genus. This is
the first record of the species fromGRSM, North Carolina, and the south-
ern Appalachian Mountains. Additional
GRSM specimens exist in the PurdueEntomology Research Collection
[PERC]: NC: Swain Co., Oconaluftee
River, 35^31 '59"N, 83°18'08"W, 15-V-
2001, C. D. & R. P. Randolph, L. M.Jacobus, one nymph. TN: Blount Co.,
Middle Prong, 35°38'30"N, 83°4r25"W,18-V-2001, C. D. & R. P. Randolph, L.
M. Jacobus, one nymph; Cocke Co.,
Cosby Creek, 35°46'59"N, 83"13'06"W,
17-V-2001, C. D. & R. P. Randolph, L.
M. Jacobus, one nymph; Sevier Co.,
Jakes Creek, 35°38'44"N, 83°35'02"W,
12-VI-2003, J. M. Webb, L. M. Jacobus,
one nymph.
Baetidae
Acerpenna macdumioughi (Ide).—Twospecimens were collected from ShopCreek and Twentymile Creek at camp-ground 93. These represent new GRSMand Tennessee records. Morihara andMcCafferty (1979) listed this species
from North Carolina, but they provided
no actual record data (sensu McCafferty
2001).
Caenidae
Caenis inacafferti Provonsha.—This
species was first reported from GRSMas Caenis nr. macafferti by DeWalt andHeinold (2005), based on adult speci-
mens from lower Abrams Creek. Wenow have larvae from Shop Creek andAbrams Creek Campground and one
reared adult of each sex from the latter
location. The larvae might be identified
as C tardata McDunnough when using
the key provided by Provonsha (1990);
they have the operculate second gill
uniformly brown and hind tarsal fimbri-
ate spurs numbering 12. However, the Y-ridge diverges in the anterior half of the
operculate gill, not in the posterior as in
C. tardata. Adults will key (Provonsha
1990) to near C. macafferti, with fore-
wing vein ICuAl forked from CuA2 just
distad of the CuAl-CuP crossvein; ab-
dominal terga 1-10 shaded blackish
brown and terga nine and 10 lacking
triads of black dots; each egg has one
polar cap. However, they lack the fleshy
protuberance characteristic of C waca/-
/^rrz (Provonsha 1990). Provonsha (1990)
lists it from Florida to New York, with
a disjunct population in the OzarkMountains of Arkansas. This is a newNorth Carolina state record.
Ephemeridae
Hexagenia limbata (Serville).—Three
specimens were collected from DeepCreek and Shop Creek. The species has
a transcontinental distribution on NorthAmerica (McCafferty 1994) including
North Carolina Pescador et al. (1999)
and Tennessee (Long and Kondratieff
VOLUME 109, NUMBER 1 149
200-
'^ 150
£ 100
aHPh
8
Stream Reaches
12
Fig. 2. Species accumulation curve for Ephemeroptera, Plecoptera, and Trichoptera species richness
from stream reaches in southwestern Great Smoky Mountains National Park, Tennessee and NorthCarolina. Solid line is predicted line-of-best-fit, dashed lines are one standard deviation out as
confidence intervals.
1996). It was reported by Spieth (1941)
from Bryson City, North Carolina, just
outside of GRSM, but this is the first
published record from within GRSM.Heptageniidae
Epeorus fragilis (Morgan).—A single
female subimago was collected fromDeep Creek. A male adult was collected
from a nearby drainage in GRSM: NC:Haywood Co., Trib. Hemphill Creek,
5 km WNW Jonathan, 35°34'35"N,
83°04'22"W, 31-V-2003, R. E. DeWalt,
Catalogue # INHS 9961. Authors LMJand WPM report additional specimens
from several streams elsewhere in
GRSM—NC: Haywood Co., Big Creek
at Big Creek picnic area, 35'45'05"N,
83°06'31"W, 12-VI-2003, J. M. Webb, L.
M. Jacobus, one nymph. TN: Sevier
Co.,Tributary Little River, 35°38'45"N,
83°35'03"W, 16-V-2001, C. D. & R. P.
Randolph, L. M. Jacobus, 4 nymphs;
Sevier Co., Injun Creek at Green-
brier Ranger Station, 35°43'49"N, 83°
24'22"W, 7-XII-2001, J. F. MacDonald,L. M. Jacobus, 4 nymphs [PERC]. Thespecies has been reported from WestVirginia (Faulkner and Tarter 1977),
Virginia (Kondratieff and Voshell 1983),
and New York (Jacobus and McCa-fferty 2001). Our data represent newGRSM, North Carolina, and Tennessee
records.
Nixe spinosa (Traver).—A single spec-
imen was taken from Deep Creek. It is
known from North Carolina (Pescador
et al. 1999) and the Abrams Creekdrainage of GRSM (DeWalt and Hei-
nold 2005). The species is seldom re-
ported, perhaps because the larva is
unknown. However, LMJ and WPMnote that adults have been collected
commonly at lights elsewhere in GRSM,especially from some of the more eastern
drainages (Jacobus and McCafferty, un-
published data).
150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
SHOP
TABCAT
EAGLE90
DEEP
EAGLE89
EKAN
HAZEL
HAMMERSEEP
CATTAIL
TWENTY93
TWENTY92
PROCTOR
GUNNA
3
0.0 1.2 2.3 3.5 4.6
Distance
Fig. 3. Unweighted pair-group method cluster analysis of Ephemeroptera, Plecoptera, andTrichoptera assemblages in stream reaches in southwestern Great Smoky Mountains National Park,
Tennessee, and North Carolina.
Rhithrogena fasciata Traver.—A large
population was taken at Eagle Creek at
campground 90. Traver (1933) listed it
from several western North Carolina
sites, including Waynesville in HaywoodCounty, adjacent to GRSM. It has been
reported also from South Carolina (Pes-
cador et al. 1999) and Georgia (Berner
1977). The species has been collected
from one other GRSM location: NC:Swain Co., Bradley Fork at Smokemont,24-IX-2002, L. M. & P.D. Jacobus, onenymph [PERC]. These are new records
for GRSM.Steiiacron pallidum (Traver).—Sixteen
specimens were collected from four loca-
tions in Tennessee and North Carolina.
Lewis (1974) reported it as restricted to
the North Carolina mountains. Subse-
quently, it was reported from SouthCarolina (Morse et al. 1989), New York
(Jacobus and McCafferty 2001), andTennessee (DeWalt and Heinold 2005)
in GRSM. Authors LMJ and WPM have
collections from widely distributed loca-
tions in the GRSM.Maccaffertiuni carlsoni Lewis.—Seven
specimens were collected as part of this
study from three reaches in Tennessee
and North Carolina, representing newGRSM and Tennessee records. Morse et
al. (1989) previously reported M. carlsoni
from North Carolina. Additional speci-
mens residing in PERC include: TN:Sevier Co., LeConte Creek, 35°40'34"N,
83°29'16'W, 16-V-2001, C. D. & R. P.
Randolph, L. M. Jacobus, one nymph;Sevier Co., Tributary LeConte Creek,
35°40'41"N, 83 28'55'W, 3-XII-2001, C.
D. & R. P. Randolph, L. M. Jacobus,
one nymph; Blount Co. Tributary An-thony Creek, 35°35'06"N, 83°45'29'W,
VOLUME 109, NUMBER 1 151
19-V-2001, C. D. & R. P. Randolph, L.
M. Jacobus, one nymph.Leptophlebiidae
Paraleptophlebia guttata (McDunno-ugh).—Single specimens were taken fromfour locations in North Carolina andTennessee. Pescador et al. (1999) listed it
from North and South Carolina andLong and Kondratieff (1996) listed it
from several Tennessee counties, includ-
ing Sevier, but neither provide specific
locality records (sensu McCafferty 2001),
so it is not possible to know if these
represent GRSM records. LMJ andWPM note that this species is generally
widespread in GRSM, and that it has
been collected from habitats ranging
from spring seeps to large streams. Theseare the first confirmed records of the
species from GRSM.Paraleptophlebia mollis (Eaton).
—
Four specimens were collected fromDeep Creek. Pescador et al. (1999)
recorded it from North and SouthCarolina and Long and Kondratieff
(1996) reported it only from Sevier
County in Tennessee. Neither work pro-
vided specific locality data. This is the
first confirmation of its presence in
GRSM.Neoephemeridae
Neoephemera purpurea (Traver).—Onelarva was collected from Deep Creek as
part of this study. It is noted here due it
apparent rarity in GRSM. Elsewhere, it
is relatively widespread, being knownfrom Florida, Georgia, Kentucky, NorthCarolina, South Carolina, Tennessee,
Virginia, and West Virginia (Traver
1935; Faulkner and Tarter 1977; Markeret al. 1979; Bae and McCafferty 1998).
Siphlonuridae
Siphlonurus typicus (Eaton).—Onespecimen was collected from Deep Creek
as part of this study. DeWalt andHeinold (2005) reported a single male
from Abrams Creek. It is known from
scattered locations in the Midwest, the
northeastern states and Canada (Provon-
sha and McCafferty 1982; Randolph andMcCafferty 1998), and from New York(Jacobus and McCafferty 2001). This is
a new North Carolina state record.
Plecoptera
Perlodidae
Oconoperla innubila (Needham andClaassen).—A male and female were
taken from Proctor Branch. Stark andStewart (1982) proposed the genus anddescribed a new species Oconoperla
weaveri. Stark (1985) later moved Yugus
innubilus (Needham and Claassen) to
Oconoperla, and synonomized O. weaveri
with it. This is a rare species from Northand South Carolina and Tennessee. It is
known from a total of seven locations,
including Clingman's Dome, GRSM(Stark 1985; Stark and Stewart 1982).
Trichoptera
Glossomatidae
Agapetus iridis Ross.—Three males
were taken from Hammer Branch. Un-zicker et al. (1982) reported it frommountain and piedmont ecoregions of
North and South Carolina. It is knownfrom two counties in Tennessee (Etnier
et al. 1998) and is a new GRSM record.
Leptoceridae
Ceraclea diluta (Hagen).—Several spe-
cimens were taken from Deep Creek. It
was reported from the North Carolina
coastal plain by Unzicker et al. (1982)
and several east-central counties of
Tennessee Etnier et al. (1998). It is
a new GRSM record.
Triaenodes taenius Ross.—A single
male was collected from Tabcat Creek.
It was described at the GRSM doorstep
by Ross (1938) and was confirmed for
GRSM by DeWalt and Heinold (2005).
Morse (unpubl. data) reported it fromthe Ravensford area of GRSM. Etnier et
al. (1998) provided one Cocke Co. record
adjacent to GRSM.Molannidae
Molanna ulmerina Navas.—Twelvespecimens were taken from Shop Creek.
DeWalt and Heinold (2005) reported it
152 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
from low elevation Abrams Creek
reaches of GRSM. It is known fromCumberland County in Tennessee (Et-
nier et al. 1998).
Polycentropodidae
Nyctiophylax denningi Morse.—Thir-
teen males were taken from Shop andTabcat creeks. Armitage and Hamilton
(1990) listed it from Alabama, Georgia,
Mississippi, South Carolina, and Ten-
nessee (the latter also by Etnier et al.
1998). It is a new GRSM record.
Sericostomatidae
Agarodes tetron (Ross) or grisea
Banks.—One female was taken fromDeep Creek. Ross and Scott (1974) prov-
ided a key to Agarodes, but could not
separate females of these species. Agar-
odes tetron has been reported from un-
specified Tennessee locations adjacent to
GRSM (Etnier et al. 1998) and fromRavensford in North Carolina by Morse(unpubl. data). This is a new generic
GRSM record.
Discussion
This inventory produced a total of 169
EPT species, four more than was foundby DeWah and Heinold (2005) in their
examination of the Abrams Creek drain-
age in GRSM. Sorensen's quotient of
similarity between these two data sets
demonstrated a 61% overlap in species
composition.
This study produced many more may-fly species, especially in the families
Heptageniidae, Ephemerellidae, and Bae-
tidae than did the Abrams Creek work.
Among the latter two families, additional
handpicking of larvae may have helped to
increase our recognition of species rich-
ness. Among stoneflies, Perlidae were not
as rich as in Abrams Creek, the majority
being lost in the genus Perlesta. Addi-tionally, caddisfly richness was much less
than for Abrams Creek, with major losses
among the Hydroptilidae and Leptocer-
idae. The lack of a large, warm, placid
stream, like lower Abrams Creek, is
probably the cause of the difficiency in
these two families.
The current study produced significant
regional records for the mayflies Amele-
tus tertius and Epeorus fragilis. Addi-
tionally, a number of first reports of
Ephemeroptera species are provided for
North Carolina (four), for Tennessee
(five), and for GRSM (seven). Fournew GRSM records of Trichoptera are
included also.
Species accummulation curves pre-
dicted that potentially 80 to 120 moreEPT species will be collected from this
region of GRSM. There are still manydrainages north of Lake Fontana that
have been poorly studied, and it is likely
that other areas will yield additional
species. For instance, further inventories
of EPT species in the highest elevations of
GRSM will yield records representing
significant range extensions for somespecies (E. Fleek, pers. comm.). Investiga-
tions of remote, higher elevation streams
in the southern Appalachians, in general,
are needed to assess the biological di-
versity of this region. For example, recent
sampling from remote stream reaches in
southern North Carolina revealed a genus
previously unreported from North Amer-ica (Waltz 2002).
Acknowledgments
We thank DLIA for funding and C.
Parker for guidance in picking locations.
DLIA employee J. Hilten arranged for
housing in GRSM, and K. Langdon and
the National Park Service have supported
our efforts and requests for permits.
Additionally, S. Johnson, J. Burbank, J.
Lowe, S. Higdon have helped pack in
heavy batteries and spent long evenings
with the ultraviolet lights in the back
country. M. Meyer, Purdue University,
identified most mayfly specimens, and A.
Provonsha, Purdue University, identified
Caenis macafferti. Finally, the efforts of
B. D. Heinold were instrumental in
collecting specimens for this study.
VOLUME 109, NUMBER 1 153
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